Mechanisms of Sound Localization in Mammals. B. Grothe, M. Pecka, D. McAlpine (2010). Presented by Alex R. Daniel M.

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1 Mechanisms of Sound Localization in Mammals B. Grothe, M. Pecka, D. McAlpine (2010). Presented by Alex R. Daniel M.

2 Introduction This paper seeks to demonstrate three ways in which our understanding of sound localization in mammals has changed with recent studies.

3 Introduction: The Three 1. Source location is represented by a bilateral population of broadly tuned spatial channels, not a topographic space-map. 2. ITD is not processed by exclusively excitatory delay lines in mammals 3. Binaural mammalian audition is not as hard wired as previously thought, and can dynamically adjust its tuning properties

4 Background - ITD - ILD - HRTF

5

6

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8 Acoustic Cues for Sound Localization in Mammals, Part A Phenomena investigated since mid 19th century Rayleigh (1907) confirmed research done by Thompson (1882) to create a duplex theory of sound localization: ILDs high-frequency noise localization ITDs low-frequency noise localization

9 Acoustic Cues for Sound Localization in Mammals, Part A <Cone of confusion>

10 Acoustic Cues for Sound Localization in Mammals, Part A Head-related transfer function (HRTF): Describes spectral modification generated by pinna and concha of outer ear. Birds and mammals can adapt to altered cues, as in the concha mold experiment: Ear mold changes the HRTF cues. After several weeks, humans adapt When molds removed, no residual effects. Learned behavior?

11 Acoustic Cues for Sound Localization in Mammals, Part B: Human Localization Humans have remarkable spatial hearing capabilities Accuracy within 1-2 degrees in angular location of a sound source Thresholds of 10 µs ITDs and 1-2 db ILDs in earphone studies Significance of ITD: action potentials take milliseconds, so 10 µs is impressive.

12 Acoustic Cues for Sound Localization in Mammals, Part B: Human Localization Steven & Newman (1934) investigated absolute localization in humans: Performance was best below ~2kHz and above 5kHz, large inaccuracies in between. Suggests incomplete representation of either cue over this range

13 Acoustic Cues for Sound Localization in Mammals, Part B: Human Localization Range of ITDs found in psychophysical studies using headphones: Range roughly consistent around frequencies below 1.5 khz Set by physiological range ± 700 µs in humans but up to ± 3,000 µs. (ITDs longer than physiological range found when multiple sources of sound interact & in acoustic reflections.

14 Neural Mechanisms in mammals

15

16 Neural mechanisms in mammal - Central auditory system consists of different ascending, segregated pathways - First stop up is the ventral cochlear nucleus. - Bushy cells respond to fine structure of sounds, are critical for binaural and project to the superior olivary complex, where ITD and ILD are processed.

17 Neural mechanisms in mammal - Central auditory system consists of different ascending, segregated pathways - First stop up is the ventral cochlear nucleus. - Monaural cues pathways project onto lateral lemniscus and contralateral inferior colliculus.

18 Neural mechanisms in mammal - IC is a necessary station for all pathways - IC is highly convergent, and it seems that all acoustic cues have already been processed

19 Neural mechanisms: vertical localization - Dorsal Cochlear Nucleus seems specialized for processing spectral cues; some spectral cues seem to be processed in IC as well. - Type IV neurons in DCN are sensitive to a circuit in the DCN, and show an island of near-threshold activity immediately around their characteristic frequency, and have a large CIA at higher sound intensity

20

21 Neural mechanisms: vertical localization - Type IV neurons get their functionality from convergence of excitatory synapses with auditory nerve fibers, and inhibitory from type II DCN neurons. - Type - II neurons are thought to respond to Onset-C inhibition. - This makes TIV neurons sensitive, inhibitorily, to acoustic notches.

22 Neural mechanisms: vertical localization - Type IV neurons project to type O neurons. - Type O neurons feature a frequency-versusintensity response area. - Respond similarly to tivs for pure tones, but have essentially the opposite response, excitatory, across the notch-center map. - tiv to to interneurons might be specialized

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